MYCOTAXON Volume 110, pp. 283–287 October–December 2009

Crepidotus subfulviceps comb. nov., a stipitate from temperate North America and Europe

M. Catherine Aime1, Jordi Vila2 & Pierre-Arthur Moreau3 [email protected] Department of Plant Pathology and Crop Physiology, Louisiana State University Agricultural Center, Baton Rouge, LA 70803 USA 2Rector Ubach, 53, ático 2a, E-08021 Barcelona SPAIN 3Laboratoire de Botanique, Faculté des Sciences Pharmaceutiques et Biologiques 3 rue du Professeur Laguesse, B.P. 83, F-59006 Lille Cedex FRANCE

Abstract — Nuclear rDNA sequence analyses indicate that Tubaria decurrens, a Pan- American species recently discovered in Spain, belongs in the Crepidotus. This is the fourth reported centrally stipitate species in the otherwise predominantly pleurotoid genus Crepidotus. The new combination Crepidotus subfulviceps is proposed. Key words — , , fungal systematics, Melanomphalia

Introduction TheCrepidotaceae (, Agaricales) s. Singer (1986) contains several genera of centrally stipitate that have since been excluded from the family based on nuclear rDNA sequence analyses and type studies (Aime et al. 2005). However, it was concluded that while the type species of Tubaria (W.G. Sm.) Gillet and Melanomphalia M.P. Christ. did not belong in Crepidotaceae, several other taxa currently allied within those genera might be more closely related to Crepidotus than to the types of the former two genera, warranting further study (Aime et al. 2005). The genusTubaria has since been shown to be polyphyletic (Matheny et al. 2007) and at least one species of Melanomphalia has been transferred to Crepidotus based on rDNA analyses (Aime et al. 2002). Recently Vila et al. (2007) found several collections of a centrally stipitate in Spain that was found to be conspecific with Tubaria decurrens, a Pan-American species originally described from Kansas. The taxon was completely described and illustrated and the authors speculated on the relationship between T. decurrens and members of the Crepidotaceae. Subsequent phylogenetic analyses of the nuclear large subunit rDNA region 284 ... Aime, Vila & Moreau confirms that this taxon is, in fact, a Crepidotus species and the new combination C. subfulviceps is proposed to accommodate it. Materials and methods A total of 19 species were chosen for analysis. Exemplar taxa, including type species from Crepidotus [type, C. mollis (Schaeff.) Staude 1857], Tubaria [type, T. furfuracea (Pers.) Gillet 1876], and Melanomphalia [type, M. nigrescens M.P. Christ. 1936], were selected from previously published Crepidotaceae nuclear large subunit rDNA (LSU) datasets (Moncalvo et al. 2000, 2002; Aime et al. 2002, 2005; Aime & Miller 2002) and from the genus Inocybe, which has been suggested as the sister group to the Crepidotaceae s.s. (Matheny et al. 2006). DNA was extracted from dried tissue from two herbarium specimens of Tubaria decurrens, placed in 2 mL Bead Solution tubes of the UltraClean Plant DNA Isolation Kit, and extracted per the manufacturer’s instructions (MoBio Laboratories, Inc., Solana Beach, CA). The first 1250 bp of the LSU were amplified and sequenced with primers LSU4-B (Aime & Phillips-Mora 2005) and LR6 (Moncalvo et al. 1995) and sequenced as previously described in Aime & Phillips-Mora (2005). Specimen vouchers are deposited in Herbarium BCN (Centre de Documentació de Biodiversitat Vegetal, Universitat de Barcelona); sequence vouchers are deposited in GenBank (http://www.ncbi.nlm.nih.gov/). For phylogenetic analyses, sequences were edited and contiguous sequences were assembled in Sequencher v.4.1.4 (Gene Codes Corp., Ann Arbor, MI). Sequence alignments were constructed by eye in Se-Al v2.0a11 (Andrew Rambaut, Dept. Zoology, University of Oxford, U.K.; http://evolve.zoo.ox.ac.uk/). Regions too variable to confidently align (a total of 27 bp in a single indel region) were excluded from the final analyses. Maximum parsimony (MP) analyses were conducted in Paup* v4.0b10 (Swofford 2002) as heuristic searches with 100 random addition replicates and TBR branch swapping. Support for the branching topologies was evaluated by bootstrap analysis derived from 1000 replicates with 10 random addition replicates each.

Taxonomy

Crepidotus subfulviceps (Murrill) Aime, Vila & P.-A. Moreau, comb. nov. Mycobank MB513297 Basionym: Omphalina subfulviceps Murrill, Lloydia 7(4): 309 (1945, “1944”). = Flammula decurrens Peck, Bull. Torrey Bot. Club 22(12): 489 (1895). ≡ Tubaria decurrens (Peck) Murrill, North American Flora 10(2): 159 (1917). ?= Tubaria omphaliopsis Singer, in Singer & Digilio, Lilloa 25: 397 (1952, ”1951”). ≡ Melanomphalia omphaliopsis (Singer) Singer, in Petersen, Evolution in the higher Basidiomycetes: 459 (1971). Material Studied: SPAIN. CATALONIA: Girona, Palafrugell, 30 Sep 2003, C. Roqué (BCN SCM B-5138), GenBank FJ947117; Prés de la Punta de la Creueta, Tarragona, 18 Oct 2005, J. Vila, X. Llimona, and L. Balcells (BCN SCM B-5144), GenBank FJ947116. Comments: Crepidotus is a genus with over 250 described species of saprotrophic pleurotoid agarics (Hesler & Smith 1965). Sequence analyses from the nuclear LSU rDNA region confirm the placement of T. decurrens within the Crepidotus subfulviceps comb. nov. ... 285

Figure 1. Mid-point rooted phylogenetic tree from MP analyses of nuclear large subunit ribosomal DNA showing position of C. subfulviceps within Crepidotus. Thickened branches are those receiving significant (>75% bootstrapping) support. genus Crepidotus (Fig. 1). The internal transcribed spacer region of the nuclear rDNA was also sequenced and analyzed with the same result (data not shown). Thus, the placement ofT. decurrens within Crepidotus marks the fourth stipitate species of Crepidotus within this otherwise pleurotoid genus (Hesler & Smith 1965, Aime et al. 2002). Phylogenetic analyses show that three of the known species of stipitate Crepidotus—C. nyssicola (Murrill) Singer 1973, C. thermophilus (Singer) Aime et al. 2002, and C. subfulviceps—do not form a monophyletic group (Fig. 1) 286 ... Aime, Vila & Moreau and therefore represent independent reversions to the stipitate habit within this genus. Crepidotus subfulviceps can be easily distinguished from the other known stipitate species of Crepidotus by its basidiospores, which are small (avg. 5 µm diam), globose and strongly echinulate in C. nyssicola versus amygdaliform in C. subfulviceps, and shorter (7.0–10.7 µm) with verrucose ornamentation in C. thermophilus versus 9.5–10.8 µm and faintly rugose in C. subfulviceps. A fourth species, Crepidotus ibericus (G. Moreno & Esteve-Rav.) Bandala et al. 2008 has recently been transferred to Crepidotus (Bandala et al. 2008) and differs from the other known stipitate species in having smooth basidiospores. The combination Crepidotus decurrens is not available for this taxon. Crepidotus decurrens States (States 1973) is a synonym for the North American C. cinnabarinus Peck 1895 (Luther & Redhead 1981), an astipitate temperate species distinguished by its scarlet-red pileus and gills that has no relation to T. decurrens. Thus, C. subfulviceps becomes the oldest available name that can be unambiguously assigned to this taxon. Crepidotus subfulviceps has been completely described and illustrated in Vila et al. 2007.

Acknowledgments The authors wish to thank Jean Lodge and Tim Baroni, as well as the Editor-in-Chief and Nomenclature Editor of Mycotaxon, for their expert review of this manuscript.

Literature cited

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